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A Review of the Cholevinae from the Island of Borneo (Coleoptera, Leiodidae)

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A Review of the Cholevinae from the Island of Borneo (Coleoptera, Leiodidae) A peer-reviewed open-access journal ZooKeys 777: 57–108A review (2018) of the Cholevinae from the island of Borneo (Coleoptera, Leiodidae) 57 doi: 10.3897/zookeys.777.23212 RESEARCH ARTICLE http://zookeys.pensoft.net Launched to accelerate biodiversity research A review of the Cholevinae from the island of Borneo (Coleoptera, Leiodidae) Menno Schilthuizen1,2,3,4, Michel Perreau5, Iva Njunjić1,3,6 1 Naturalis Biodiversity Center, Darwinweg 2, 2333CR Leiden, The Netherlands2 Institute for Biology Leiden, Leiden University, Sylviusweg 72, 2333BE Leiden, The Netherlands3 Taxon Expeditions, Rembrandtstraat 20, 2311 VW Leiden, The Netherlands 4 Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah, Jalan UMS, Kota Kinabalu, Malaysia 5 IUT Paris Diderot, Université Paris Diderot, Sorbonne Paris cité, case 7139, 5 rue Thomas Mann, 75205 Paris cedex 13, France6 Institut de Systématique, Évolution et Biodiversité, Muséum National d’Histoire naturelle, Sorbonne Universités, Paris, France Corresponding author: Menno Schilthuizen ([email protected]) Academic editor: B. Guéorguiev | Received 23 December 2017 | Accepted 5 June 2018 | Published 30 July 2018 http://zoobank.org/D9F35364-3DCD-4BA6-B70D-62FB275DEB1B Citation: Schilthuizen M, Perreau M, Njunjić I (2018) A review of the Cholevinae from the island of Borneo (Coleoptera, Leiodidae). ZooKeys 777: 57–108. https://doi.org/10.3897/zookeys.777.23212 Abstract The available knowledge of the round fungus beetle subfamily Cholevinae (Leiodidae) from the island of Borneo is reviewed, and the results of newly studied material presented. The currently known 30 species (of which 14 are newly described herein) represent the genera Micronemadus (one species), Catops (one species), Baryodirus (one species), Ptomaphaginus (14 species), and Ptomaphaminus (13 species). The following new species are described: Micronemadus sondaicus Schilthuizen & Perreau, sp. n., Ptomaphaginus grandis Schilthuizen & Perreau, sp. n., P. louis Schilthuizen & Perreau, sp. n., P. muluensis Schilthuizen & Perreau, sp. n., and P. isabellarossellini Schilthuizen, Njunjić & Perreau, sp. n., and Ptomaphaminus kinabatanganensis Njunjić, Schilthuizen & Perreau, sp. n., P. testaceus Schilthuizen & Perreau, sp. n., P. nanus Schilthuizen & Perreau, sp. n., P. marshalli Schilthuizen & Perreau, sp. n., P. hanskii Schilthuizen & Perreau, sp. n., P. sarawacensis Schilthuizen & Perreau, sp. n., P. layangensis Schilthuizen & Perreau, sp. n., P. microphallus Schilthuizen & Perreau, sp. n., and P. alabensis Schilthuizen & Perreau, sp. n. It is expected that the cholevine biodiversity of Borneo is still far from completely known. Nonetheless, provisional identification keys to all species known so far are presented. Keywords caves, Indonesia, Malaysia, scavenging beetles, taxonomy Copyright Menno Schilthuizen et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 58 Menno Schilthuizen et al. / ZooKeys 777: 57–108 (2018) Introduction Borneo is, after New Guinea, the second-largest tropical island of the world. It has never been strongly isolated, having formed part of a larger land mass, known as Sun- daland, during marine transgressions in the Pleistocene (Hall and Holloway 1998). Sundaland comprises the present-day Malay Peninsula, the islands of Sumatra, Java, and Borneo, as well as the shallow seas in between. Recent paleoclimatic modelling (Raes et al. 2014) suggests that even during cooler periods of marine regression, ever- green wet forests dominated the area that now comprises Borneo. This, and the fact that the island, because of its mountainous character (with Gunung Kinabalu reaching almost 4,100 m) contains a great variety of habitats, has generated and maintained a rich biodiversity, not least in its soil and litter-dwelling invertebrate fauna (e.g., Hanski and Hammond 1986, Rahman et al. 2002, Liew et al. 2010). The beetle subfamily Cholevinae (Coleoptera, Staphylinoidea, Leiodidae) consists mostly of small, soil-dwelling scavengers, well represented in the litter fauna of all tropical regions. Szymczakowski (1964) provided the first overview of the Cholevinae of South Asia, but the large number of new species described since then (Perreau 2000, and unpublished checklists) mean that this work is severely outdated by now. In recent years, several semi-comprehensive studies have appeared, either limited to a certain region (e.g., Wang and Zhou 2015) or to a certain habitat (e.g., Perreau 2009). In this paper, we provide an overview of the species of Cholevinae currently known from the island of Borneo. By necessity, this is a very preliminary overview, since it is based on comparatively little information. Jeannel (1936) mentions only one species from Borneo. Szymczakowski (1961) mentions one more, and Peck (1981) and Perreau (2000) describe two more. In 2008, we (Schilthuizen and Perreau 2008) described seven new species and two new records from Borneo, bringing the total cholevine fauna to 13 spe- cies. However, more recent work (e.g., Merckx et al. 2015), as well as study of existing material in the Natural History Museum (London), deriving from the 1978 Mulu expedi- tion (Hanski 1983, Hanski and Hammond 1986), and in Naturalis Biodiversity Center (Leiden), have revealed many additional new species. In our opinion, the 30 species that we recognize in the present paper form a sufficient basis to produce a first overview of our current, but doubtlessly still very incomplete, knowledge of the cholevine fauna of Borneo. We provide a brief description for all previously described genera and species, and more extensive descriptions for newly-described species, as well as differential diagnoses for new species that have close congeners in Borneo. Where available, we also refer to DNA sequences in the Barcode of Life Database (BOLD, http://boldystems.org) and to so-called Barcode Index Numbers (BINs; Ratnasingham and Hebert 2013). We also give preliminary identification keys. However, given the fact that this overview is probably still far from complete, these keys should be used with caution: any sample collected in Borneo is likely to contain previously unrecognised species, and we hope this paper will stimulate further taxonomic and faunistic work. A review of the Cholevinae from the island of Borneo (Coleoptera, Leiodidae) 59 The collection abbreviations used in the lists of examined material are as follows: BORN Borneensis Collection, Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah, Kota Kinabalu, Malaysia; FRCS Forest Research Centre, Sepilok, Malaysia; HNHM Hungarian Natural History Museum, Budapest, Hungary; JRUC collection of Jan Růžička, Prague, Czech Republic; MHNG Muséum d’Histoire Naturelle de Genève, Switzerland; NHMUK Natural History Museum, London, UK (formerly British Museum (Natural History)); RMNH Naturalis Biodiversity Center, Leiden, The Netherlands (comprising the collections of the Rijksmuseum van Natuurlijke Historie, Leiden and the Zoölogisch Museum, Amsterdam). TXEX Taxon Expeditions, Leiden, The Netherlands. CMPR Collection Michel Perreau, Paris. Male genitalia were mounted in Euparal after dissection and dehydration in ethanol 95%. Female genitalia were cleared in hot KOH 10% and stained with Azoblack before mounting in DMHF (2,5-DiHydroxyMethylFurane). Photographs of genitalia were taken on a Leitz Diaplan microscope using a Spot Insight IN1820 or a Leica MC170HD camera. High-resolution photonic pictures of external mor- phological details (Figure 14a–e) were taken with a Keyence VHX5000 microscope and a VH-Z250T lens. The outline of the map was built from SRTM3 data (Shuttle Radar Topography Mission) of NASA with the software QGIS. They were com- pleted by the global administrative areas database GDAM (www.gdam.org) and the hydrographic networks of Natural Earth (http://www.naturalearthdata.com/). All images and descriptions will be uploaded to the Cholevidae of the World website (http://cholevidae.myspecies.info/). Systematics Tribe: ANEMADINI Hatch Genus: Micronemadus Jeannel, 1936 Description. Small, 1.4–2.6 mm. Oval habitus, with the apex of the elytra rounded. Head and pronotum punctate, elytra with transverse strigae. Antenna with antennomer- es 4 and 5 very short and wide. Mesosternum with a longitudinal mesoventral process. The four first protarsomeres and the first mesotarsomere dilated in the male. Aedeagus: median lobe triangular, parameres longer than the median lobe, and bent inward. 60 Menno Schilthuizen et al. / ZooKeys 777: 57–108 (2018) Micronemadus sondaicus Schilthuizen & Perreau, sp. n. http://zoobank.org/8AA2A703-76B4-4DCE-BFD3-8E19943EA8BA Figure 1a, c–f Material. Holotype: Malaysia, Sabah, Crocker Range Park, Inobong, 5°51.265'N, 116°08.363'E, 500 m elev., 21–23.ix.2012 (leg. M. Schilthuizen, Crocker Range / Kinabalu Expedition, RMNH.INS.555641), male. Paratypes: Sabah. Crocker Range Park, Inobong, 5°51.3'N, 116°08.4'E, 500 m elev., 21–23.ix.2012 (leg. M. Schilth- uizen, Crocker Range / Kinabalu Expedition, RMNH.INS.555640, 555642), 2 in- dividuals; Crocker Range, Gunung Alab, xii.2009 (leg. M. Schilthuizen, RMNH. INS.63291, 63310, 63295), 3 individuals; Crocker Range, along the road from Kota Kinabalu to Tambunan, near Rafflesia Park, 5°46.4'N, 116°20.8'E, 1350 m elev., baited pitfall trap, 2001 (leg. M. Schilthuizen,
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